Pseudoliodini (Coleoptera: Leiodidae: Leiodinae) of New ZealandRichard A. B. Leschen New Zealand Arthropod Collection, Landcare Research, Private Bag 92170, 120 Mt Albert Road,Auckland, New Zealand, email: LeschenR@landcare.cri.nz

AbstractThe New Zealand Pseudoliodini include two

genera and three species that are described as new:Zelodes n. gen. (Type species: Z. kuscheli n. sp.; andZ. minutus n. sp.) and Colenisia zelandica n. sp.The species of Zelodes are recognized by several distinctive characters that are unique forthe tribe: eyes absent; patch of peglike setae present on labrum; mesosternum with a triangularligulate process; ventrites I and II connate;tarsomeres not sexually dimorphic and 4-4-4;tenent setae present on pro- and mesotarsomeres1 - 3. Notes on the identification of New Zealand leiodids and a discussion on characters associatedwith hind wing reduction in Leiodidae are included.

Key words: Zelodes, Colenisia, mycophagy,wingloss, new species.

IntroductionA significant portion of beetles occurring in leaf

litter are species in the superfamily Staphylinoideawhich includes the family Leiodidae. Species ofLeiodidae are biologically diverse and feed onfungi, are saprophagous, or feed on the decay fluids or microorganisms in carrion. Because manyspecies of Cholevinae and Leiodinae are associatedwith different microhabitats and are generally distributed throughout forests, Leiodidae aremodel candidates as bioindicators of forest healthand of precise classes of forest communities (Topp& Engler 1980; Chandler & Peck 1992; Ruzicka1994; Carlton & Robison 1999). For example,selective cutting in eastern North American oldgrowth forests of different ages showed a differ-ence in leiodid species abundance while speciesrichness was maintained (Chandler & Peck 1992).Correlations between leiodid communities andtypes of ecosystems have not yet been determinedin New Zealand. The leiodid fauna here is composed of winged and flightless species, and arigorous biodiversity study using various trappingmethods may show some interesting patterns ofseasonal distribution and habitat selection betweenthe two morphological classes. One hindrance to

understanding underlying ecological patterns andphylogenetic relationships is the poor taxonomicknowledge of the family in New Zealand and thenatural history of each species.

Studies on groups such as the Camiarinae wouldrequire a comprehensive study of numerous undescribed taxa to identify monophyletic highertaxa.The family-group classification reviewed byNewton (1998) provides an appropriate background for study of the family in New Zealandwhich consists of species in four subfamilies(Camiarinae, Cholevinae, Coloninae, andLeiodinae).They range in size from about 1.0 - 4.0mm. Most are round or oval in outline while manymembers of the tribe Camiarini (Camiarinae) maybe somewhat biconvex.The largest and mostdiverse taxon is the subfamily Camiarinae whichhas 12 genera and approximately thirty describedspecies.These small Coleoptera can be identifiedusing one or both of the family-level keys inKlimaszewski and Watt (1997) with cautionbecause some of the couplets are vague and not allgenera will key to Leiodidae.

In this paper the species of Pseudoliodini(Leiodinae) are reviewed and one genus (alluded toin Newton 1998), containing two species, isdescribed as new.The genus Colenisia Fauvel waspreviously recorded in New Zealand by Kuschel(1990) and the single species is formally describedherein.

Specimens examined are deposited in theEntomology Research Museum (LincolnUniversity, Canterbury, LUNZ); Field Museum ofNatural History (Chicago, FMNH); New ZealandCollection of Arthropods (Auckland, NZAC);Otago Museum (Dunedin, OMNZ); and Museumof New Zealand Te Papa Tongarewa (Wellington,MONZ).

Specimens of the taxa described in this paperwere compared to specimens of AllocolenisiaDaffner (1 sp.,Thailand), Colenis Erichson (4 spp.,North and Central America), Colenisia (3 spp.,Africa, New Caledonia), DermatohomoeusHlisnikovsky (2 spp., India), Neohydnobius Jeannel

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(1 sp., Chile), Pseudcolenis Reitter (3 spp., Borneo,Japan, Sumatra), and two undetermined genera (2 spp., Africa, Ecuador). Otherwise comparisonswere made to illustrations and descriptions in pub-lished work by Daffner (1983, 1986, 1988, 1989,1990, 1991), Peck (1998), and Wheeler (1986).

Key to the subfamilies of Leiodidae, the tribesof Leiodinae and species of Pseudoliodini

The identification of microcoleoptera can be difficult because character systems are not easilyvisible unless high magnification is used.For beginning students of Coleoptera, especiallythose who do not have access to a reference

Figs 1 - 9. (1) Head of Paracatops sp., dorsal view (scale bar =0.10 mm); (2) head of Colon sp., lateral view (scale bar = 1.0mm); (3) same, dorsal view of head (scale bar = 0.10 mm);(4) Zearagytodes maculifer (Broun), dorsal view (scale bar =

0.50 mm); (5) antennal club of Zeadolopus sp., dorsal view(scale bar = 0.10 mm); (6) antenna of Zelodes kuscheli, n. sp.,dorsal view (scale bar = 0.10 mm); (7) same, dorsal view ofhead (scale bar = 0.10 mm); (8) head of Colenisia

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collection, the keys in Klimaszewski and Watt(1997) may prove challenging. Some micro-coleoptera may be misidentified as Leiodidae;however, most New Zealand leiodids have antennomere VIII smaller than VII and IX (Fig. 6;except Colon Herbst, Fig. 2; Zearagytodes Jeannel,Fig. 4, and an undescribed genus of

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caledonica Fauvel, anterolateral view (scale bar = 0.10 mm);(9) same, prosternum, ventral view (scale bar = 0.10 mm).

Neopelatopini), and VIII may be concealed fromview as in Zeadolopus Broun (Fig. 5). Other keycharacters for Leiodidae are the relatively stronglyprojecting procoxae and an excavate hind coxa(like that shown in Fig. 22) which may differentiatethe family from similar looking Cucujoidea.

A key to the subfamilies by A. Newton is included in Klimaszewski and Watt (1997) and ismodified below (based also on Newton 1998) toinclude the Leiodinae tribes and species ofPseudoliodini occurring in New Zealand.Numbers of described genera are listed after eachfamily group, while an asterisk (*) indicates subfamilies containing undescribed genera.

1 Antennal insertions hidden in dorsal view (Fig. 8) ....................................Leiodinae; 4

- Antennal insertions visible in dorsal view (Figs 1, 2) ..............................................2

2 Head with a distinct occipital crest (Fig. 1) ...............................................Cholevinae (Mesocolon Broun, Pseudonemadus Portevin,Paracatops Portevin)

- Head without a distinct occipital crest (Figs 2, 3) ..............................................3

3 Antennomere 8 equal to 9 and 10 (Fig. 2),epistomal suture absent...................Coloninae (Colon)

- Antennomere 8 smaller than 9 and 10 (Fig. 6),or antennomere 8 equal to 9 and 10 (Fig. 4) andepistomal suture present .............Camiarinae*(Agyrtodini: Agyrtodes Portevin, ChelagyrtodesScymczakowski, Zeagyrtodes Broun, ZeagyrtomaScymczakowski, Zearagytodes; Camiarini:Baeosilpha Broun, Camiarites Jeannel, CamiarusSharp, Inocatops Broun, Zenocolon Broun;Neopelatopini: Catopsolius Sharp)

4. Antennal club compact; antennomere VIII hidden between VII and IX (Fig. 5)...................................Leiodini (Zeadolopus)

- Antennal club not compact; antennomere 8 visible between 7 and 9 (Fig. 6) ...................5

5. Epipleuron visible in lateral view; tarsal formula5-5-5 ..........................................Sogdini*(Isocolon Broun)

- Epipleuron not visible in lateral view; tarsal formula 5-4-4 or 4-4-4 ........... Pseudoliodini 6

6. Eyes present (Fig. 8); dorsal surfaces with asparse vestiture of setae .......................Colenisia zelandica, new species

- Eyes absent (Fig. 7); dorsal surfaces without avestiture of setae .............Zelodes, new genus; 7

7.Width of prothorax equal to length of elytra(Fig. 27) .......................Z. kuscheli, new species

- Width of prothorax shorter than length of elytra(Fig. 28) .....................Z. minutus, new species

Tribe Pseudoliodini Portevin, 1926This is a relatively small group of Leiodinae with

11 genera and about 120 species described worldwide, though most faunas of the world requirestudy. Newton (1998) characterised this groupbased on several characters in his key to theLeiodinae tribes and in the diagnosis for the tribe.He provided evidence that Pseudoliodini is relatedto the tribes Agathidiini and Scotocryptini,

although the exact relationships of these to oneanother are unknown as they are grouped into anunresolved trichotomy in his proposed phylogeny.

Pseudoliodini are collected most frequently bysifting leaf litter, although it is not surpising thatColenisia zelandica may be associated withBasidiomycetes because many leiodids aremycophagous.Three genera and several species ofPseudoliodini (including Australian species ofColenisia) have been collected from variousBasidiomycetes (Newton 1984). I have seen norecords for larval Pseudoliodini from fungi,despite the relatively common occurrence of somespecies in mushrooms in North America (Leschen1988, Peck 1998) (Note that Newton (1984) hasreared larvae of North American Colenis from sapflows). Similar habits may be shared for thespecies of Colenisia described below because it was collected three times from Basidiomycetes:Boletus sp. (Boletaceae), Coltricia cinnamomea

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Figs 10 - 16. Zelodes kuscheli, n. sp. (10 - 15): (10) labrum,anterior view same, (scale bar = 10.00 um); (11) same, detailof sensory setae (scale bar = 5.00 um); (12) left maxilla, ven-tral view (scale bar = 0.10 mm); (13) mentum and labium,ventral view (scale bar = 0.10 mm); (14) right maxilla

showing detail of lacinea and galea, ventral view (scale bar =0.05 mm); (15) right mandible, ventral view (scale bar =0.10 mm); (16) meso-, meta-, and abdominal sterna ofColenisia caledonica, ventral view (scale bar = 0.10 mm).

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(Polyporaceae), and Amanita muscaria(Amanitaceae) that was infected with Hyphomyces(Ascomycetes). It is curious that specimens of ourColenisia were collected from Hyphomyces-infect-ed Amanita because many of the records of Colenisspp. recorded in Leschen (1988) were also fromparasitised mushrooms.The gut of one paratype ofColenisia zelandica was packed with spores ofColtricia cinnamomea.

Genus Colenisia Fauvel (Figs 8, 9, 16, 26, 29, 34-37)

Colenisia zelandica, n. sp.(Figs 26, 29, 34-37)

Description: Length 1.37 mm (1.30 1.45; n =6), width 1.00 (0.87 1.12), depth 0.44 (0.370.50). Colour of body dark and light yellowbrown; head in most specimens, elytra, antennalclub and disk of mesosternum darker. Relativelengths of antennomeres3:5:3:1.2:1:1.1:3:1:3:2.1:5. Head, pronotum andelytron with well developed transverse strigulae.Epistomal suture straight and weakly impressed.Male tenent setae present on protarsomeres 1-3.Aedeagus 2 times longer than wide, aedeagal apexevenly rounded or slightly acute; parameres reaching to apical 1/5 of aedeagus, apex bisetoseand arising from bifurcate apex, setae of subequalsizes; internal sac complex. Spermatheca with distal chamber slightly longer than globular proximal chamber. Gonocoxite with three elongate setae; stylus with three subequal setae.

Etymology: Based on the distribution of thespecies in New Zealand.Remarks: Colenisia zelandica is known from thenorthern portion of the North Island. Based onaedeagal characters described by Daffner (1986,1989), C. zelandica may be related to some of theColenisia species of New Caledonia and New Guinea. In some of these species and C. zelandica, the apical setae of the parameres aresubequal in length. C. zelandica differs from all thespecies by having two anterior lobes arising fromthe base of the internal sac.The described Colenisiaspecies from Australia have a very short inside (ordorsal) apical seta, half or less the length of theouter seta. Colenisia zelandica and C. serica Daffnershare a pair of well developed symmetrical scle-rites in the middle of the internal sac, but differ inthe shape of the apex of the aedeagus.

A single teneral male specimen of the genusColenisia was collected from the Three KingsIslands and appears to be similar to C. zelandica, butwas not dissected and included in the type series.It differs externally from the specimens C. zelandica by its poorly developed transversestrigulations of the body.

While larval habits are unknown, adults are associated with fresh and decaying Basidiomycetes.Type Material:

Holotype: New Zealand, AK, Lynfield,Tropicana Drive, 10 July 1976, G. Kuschel, bushmargin (NZAC). Pa r atypes: New Zealand: 1, AK,Hunua,Waharau Reserve, 23 May 1999, R.

MONZ, NZAC, OMNZ); 1, ND,Tapotupotu Bay,30 July 1998, R. Leschen, R. Hoare leaf litterfungi, RL222, 34˚26´S, 172˚42´E (NZAC); 1, ND,Waipoua SF, 25 Nov 1980, G. Kuschel, sifted litter80/121(NZAC); 2, ND, Puketi National Forest,Nature Track, RL374, 35˚13´S 173˚47´E,ex Amanita muscaria infected by Hyphomyces,31 March 1999, R. Leschen (NZAC).

Leschen, R. Hoare leaf litter and fungi berlesate,RL412, 37˚03´S, 175˚17´E;1, same, but, Boletussp., RL413; 1, AK, Lynfield WB, 30 Sep 1980,G. Kuschel, litter (NZAC); 1, AK, KaiparaHarbour, Kaukapakapa, 23 Sep 1977, G. Kuschel,litter (NZAC); 6, ND, Ranfurly Scenic Reserve, 29July 1998, R. Leschen, R. Hoare, ex Coltricia cinnamomea, RL200, 35˚02´S, 173˚45´E (LUNZ,

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Figs 17 - 25. Zelodes kuscheli, n. sp.: (17) prosternum, ventralview (scale bar = 0.50 mm); (18) Meso- and metasternum,ventral view (scale bar = 0.10 mm); (19) same, showingdetail (scale bar = 0.10 mm); (20) fore leg of male, anterior

view (scale bar = 0.10 mm); (21) middle leg of male, anteriorview (scale bar = 0.10 mm); (22) hind leg, anterior view(scale bar = 0.50 mm); (23) detail of mesotarsus showingtenent setae of male (scale bar = 0.05 mm); (24) apex of

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Other material examined: New Zealand,TH,Great I,Tasman Valley, NZMS 260 L01 316823, 7-10.xii.1996, J.W. M. Marris, yellow pan traps inKunzea broadleaved forest (LUNZ).

Genus Zelodes, n. gen.(Figs 6, 7, 10-15, 17-25, 27, 28, 30-33, 38, 39)

Type species: Zelodes kuscheli new speciesDiagnosis: Eyes absent; patch of peglike setae

present on labrum; mesosternum with a triangularligulate process; length of metasternum subequalto width of mesocoxa; metasternum of male without setiferous sex patch; elytra with transverse striations; epipleuron completely hiddenin lateral view; wings absent; ventrites I and II connate; tarsomeres not sexually dimorphic and 4-4-4; tenent setae present on pro- and mesotarsomeres 1-3; basal lobes of parameresmeeting dorsally and fused to aedeagal body;spermatheca elongate, consisting of a single bodywith well developed cuticular rings.Description: Length about 1.4-1.7 mm.Colour dark red or yellow brown. Body complete-ly glabrous dorsally. Head transverse with 2 pairs of widely spaced pores on the vertex.Antenna about 1.7 x as long as head width, with aweak 5-segmented club; segments 7, 9, and 10with periarticular gutter, internal vesicles absent.Epistomal suture straight and very weaklyimpressed. Labrum subtransverse, apical marginwithout a shallow emargination, bearing a shortand dense patch of peglike setae. Epipharynx without a short apical furrow. Mandibles asymmetrical along inside edge, with preapicalteeth, well developed prostheca, and a strongmolar lobe at base. Labium transverse; pair of stoutsetae present. Lacinia and galea of equal widths;lacinia with midlateral spines present amongstmarginal setae; galea without field of micropores.Tentorium with or without median spine; nottuberculate. Eyes absent; field of setae present inocular area. Occipital bead narrow. Antennalgrooves absent, genal shelf rounded and not visiblein dorsal view. Prothorax without a well-developedanterior angle; prosternum relatively well devel-oped with a narrow prosternal process; ventralsurfaces of prosternal process and anterior portionof prosternum contiguous and on the same plane.Mesosternum with broad prepectus and a narrowmedian carina; a triangular ligulate process present. Metasternal process between mesocoxaedepressed anteriorly, its surface separated from andnot at same level as mesosternum. Metasternalwidth subequal to width of mesocoxae; disc notvaulted; subcoxal bead of metasternum present;pronged posterior process more or less parallelsided; male without setiferous sex patch. Length ofmedian stalk of metendosternite shorter thanlength of arm. Elytra with transverse striations;humeral region with distinct microsculpture;

male abdomen, posterior view (scale bar = 0.10 mm); (25)detail of frayed edge of ventrite VI of male, ventral view (scalebar = 10.00 um).

epipleuron completely hidden in lateral view. Hindwings absent. Abdominal ventrite VI apically frayedin male; ventrites I and II connate.Tarsal formulanot sexually dimorphic and 4-4-4; tenent setae ofmale present on pro- and mesotarsomeres 1-3.Aedeagus 3 times longer than wide and in bodycavity extending to a level of middle of metaster-num, apiculate with a squarish apex; paramereslacking apical setae, basal lobes of parameres meeting dorsally and fused to aedeagal body.Spermatheca elongate, consisting of a single bodywith well developed cuticular rings, accessory sacpresent.Etymology: A combination of parts of the wordsNew Zealand and Leiodes Latreille.Remarks: The Leiodinae tribe Pseudoliodini currently contains 11 genera (Newton 1998)including Zelodes. This genus is placed into thistribe based on the following characters (Newton1998): head without antennal grooves, procoxal

cavities closed laterally by a triangular notalprocess (Fig. 17), mesocoxae rather widely separated by a metasternal process (Figs 18, 19),tarsal formula usually not sexually dimorphic, andfree parameres of aedeagus with basodorsal lobesthat may meet dorsally (although the parameres arefused rather broadly with the base of the medianlobe as shown in Figs 32, 33). Zelodes is unlikeother pseudoliodines and can be easily distinguished from the remaining species by its lackof eyes, presence of a broad ligulate process on themesosternum, and tenent setae present on pro- and mesotarsi of the male.

Phylogenetic relationships among the genera ofPseudoliodini are basically unknown, and deter-mining the sister taxon of Zelodes is problematic.Many characters present in Zelodes are unique anddo not provide ample information for determiningits placement in the tribe. For example, theprosternum is reduced (Fig. 9) and the

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Figs 26 31. Dorsal habitus and spermathecae ofPseudoliodini: (26) Colenisia zelandica, n. sp., habitus; (27)Zelodes kuscheli, n. sp, habitus; (28) Zelodes minutus, n. sp,habitus; (29) Colenisia zelandica, n. sp., spermatheca; (30)

Zelodes kuscheli, n. sp, spermatheca; (31) Zelodes minutus, n. sp,spermatheca. (Scale bars for Figs 26 - 28 = 1.00 mm; scalebars for 29 - 31 = 0.01 mm).

26 27 28

31

29

30

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mesosternum is carinate in most species ofPseudoliodini (Fig. 16); but the prosternum is relatively well developed (Fig. 17) and the mesosternum is modified in Zelodes (Figs 18, 19).

The guts of three dissected specimens of Zelodeswere filled mostly with unidentified matter, andpossibly spores and hyphae in one specimen of Z. kuscheli.

Zelodes kuscheli, n. sp.(Figs 6, 7, 10-15, 17-25, 26, 30, 32, 33, 38)

Diagnosis: Colour of body dark red brown;pronotum about as wide as length of elytra;spermatheca with accessory sac attached proximalto spermathecal duct attachment.Description: Length 1.72 mm (1.55 - 1.85; n =9), width 1.28 (1.12 - 1.45), depth 0.63 (0.70 -0.75). Colour of body dark red brown. Headstrongly punctate, punctures separated by 1-2diameters; punctation may consist of punctures oftwo sizes in some specimens. Epistomal bead welldeveloped. Relative lengths of antennomeres5:5:4:3:3:3:4:2:5:5:7.Tentorial spine present.Pronotum with punctures lightly impressed inmost specimens, more defined at lateral margins;punctures separated by 3 diameters. Anterior edgeof furcal arms of metendosternite irregular; widthgreatest at middle. Elytron with dense punctationin anterior half; punctures separated by 1 diame-ter. Aedeagus complex with internal sac consistingof a broad anterior plate, a middle portion with aseries of smaller overlapping plates and the posterior portion membranous. Spermatheca withaccessory sac attached proximal to spermathecalduct attachment, cuticular striations dense.Gonocoxite with the two lateral setae equal inlength to single inside one.Etymology: Patronymic for Willy Kuschel, one ofthe collectors of this species and in honour of hismany contributions to the knowledge of New Zealand Coleoptera.Remarks: This species, known only from thenorthern portion of the South Island, can be distinguished from Z. minutus by its relative sizeand body shape.Type material:

Holotype: New Zealand, SD, Picton,Shakespeare Bay, 11 Aug 1969, J. McBurney, Litter69/144 (NZAC). Pa r atypes: 1, same data asholotype (NZAC); 1, SD, Port Ligar, 1000’,26 Oct 1969, F. A. Alack, Litter 69/175 (NZAC);

6, SD, Port Underwood Sdle, 1 Sep 1969,G. Kuschel, Litter 69/152 (LUNZ, OMNZ,NZAC); 1, SD, Port Underwood Saddle, 3 km SSECurious Cove, 15 Nov 1999, R. Leschen, siftingleaf litter, RL480 41˚17´S, 174˚05´E (NZAC);1, MB, Richmond Ra, Onamalutu Valley, 5 Jul1947, Leaf mould (MONZ); 1, MB, PelorusBridge, 20 Oct 1963, J. I.Townsend (FMNH);1, MB, Pelorus Bridge, 25 Sep 1967, G. Kuschel,Litter 67/220 (NZAC); 1 (slide mounted), MB, MtRobison Ridge, Kenepuru Sd., 500 m, 13 Mar1970, J. I.Townsend, litter (NZAC).

Zelodes minutus, n. sp.(Figs 28, 31, 39)

Diagnosis: Colour of body yellow brown; widthof pronotum less than length of elytra;spermatheca with accessory sac attached distal tospermathecal duct attachment.Description: Length 1.40 mm (1.40 - 1.62; n=3), width 1.04 (0.95 - 1.12), depth 0.30 (0.27 -0.35). Head with punctures separated by 1-2 diameters. Epistomal bead poorly developed.Relative lengths of antennomeres5:5:4:3:3:2:3:2:4:4:6. Pronotum lacking welldefined punctures.Tentorial spine absent. Anterioredge of furcal arms of metendosternite smooth;width greatest at base. Elytron with punctation pre-sent in anterior half; punctures separated by 1diameter. Spermatheca with accessory sac attacheddistal to spermathecal duct attachment, cuticularstriations sparse. Gonocoxite with the two lateralsetae much shorter in length than the single inside one.Etymology: Based on the Latin word minutus,meaning small.Remarks: All too little is known about thisspecies, because specific locality data are lackingand it is known only from females.Type material:

Holotype: New Zealand, taken from decaying potatoes, Coll. A. Philpott, A. E. BrookesCollection (NZAC).Pa r atypes: 4 (1 slide mount-ed and 1 with head missing), same data as holotype(NZAC).

Characters Associated with Hind WingReduction in Zelodes

Almost every group of Coleoptera includesspecies that have undergone some form of hindwing reduction and there are some excellent

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Figs 32 - 39. Terminal segments of Pseudoliodini:(32) Zelodes kuscheli, n. sp, aedeagus, ventral view;(33) same, lateral view; (34) Colenisia zelandica n. sp., aedea-gus, lateral view; (35) same, dorsal view; (36) same, close upof apex of left paramere; (37) same, dorsal view of left

gonocoxite; (38) Zelodes kuscheli, n. sp, dorsal view of leftgonocoxite; (39) Zelodes minutus, n. sp, dorsal view of leftgonocoxite. (Scale bars for Figs 32, 35, and 37 - 39 = 0.01mm; scale bar for 33 and 34 = 0.05 mm).

32 33 34 35

36

37 38 39

reviews (e.g., Roff 1990,Thayer 1992) and taxon-specific papers that document this phenom-enon (e.g., see Holloway 1963 for New ZealandLucanidae).The incidence of hind wing reductionis very high in New Zealand Coleoptera, as expected for insular faunas (Darlington 1936).Hind wing loss or reduction in Leiodidae is fairlycommon, and the phenomenon has been reviewedfor cave and soil dwelling North American speciesin the genus Ptomaphagus Illiger by Peck (1973).The hind wing may be reduced to a narrow stripretaining some of its wing veins, reduced to asmall flap that lacks veins, or completely absent asin Zelodes.

Leiodids that have undergone wing reductionmay have several characteristic modifications in therest of the body.The body shape of Zelodes is ratherordinary and not as streamlined as the body shapein many apterous cholevines - the round or oblongbody form of Zelodes species is also present inwinged pseudoliodines. In Camiarinae the bodyforms are variable, and hind wing reduction maybe associated with biconvexity (e.g., someCamiarini) which is present in other beetle families that primitively have the body outlinemore or less continuous (e.g., Languriidae;Zablotny and Leschen 1996).

Eye reduction and other characters may correlate with leiodid wing loss (Thayer 1992), insoil and cave dwelling Cholevinae andCatopocerinae, inquiline Scotocryptini, and mammal associated Platypsyllinae. Completeabsence of eyes and red coloration of Zelodes isunique in Pseudoliodini. There is a correspondingchange of the metendosternite and a shortening ofthe metasternum with the loss of flight muscula-ture in Coleoptera, and Zelodes has a relativelyshort median stalk of the metendosternite (notshown) and a short metasternum (compare Figs 16and 19). Members of Pseudoliodini typically have awell-developed median carina on the mesoster-num which is often contiguous with a vaultedmedian portion of the metasternum (Fig. 16; thisarea is poorly developed in one subgenus ofPseudcolenis). Instead of having a typical carinatemesosternum, Zelodes bears a flattened triangulateprocess between the mesocoxae (Fig. 19) and itsdevelopment may be associated with wing reduction in this genus.

A more detailed analysis of form associated withhind wing loss for New Zealand Leiodidae would

be interesting because of the variable body formspresent in the fauna. Moreover, the unusual habitsamong New Zealand taxa, such as the relativelyfast- running and flight capable species ofZearagytodes (Camiarinae) associated with polyporefungi, suggest that the different body forms maycorrelate with gait speed and performance.

AcknowledgementsI am indebted to Al Newton and Stewart Peck

for cultivating my interests in Leiodidae by supply-ing unpublished keys and catalogues, and they,along with Ivan Löbl, supplied comparative material on which this study is based.WillyKuschel helped vicariously with this projectbecause of his interests in the family that resultedin a well sorted reference collection complete withmanuscript names for undescribed genera. BirgitRhode provided excellent SEM photographs andGrace Hall helped extensively in the field andassisted with data entry. Identification of the polypore host of Colenisia zelandica was made byPeter Buchanan and habitus and line drawings weredone in part by Des Helmore and CatherineLawton. John Marris drew my attention to thespecimen of Colenisia collected from the ThreeKings Islands and Robert Hoare commented on adraft of the manuscript and also assisted in thefield. This work was made possible by funds pro-vided by FRST (contract C09617).

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